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From the Late Triassic Riograndia Assemblage Zone (Santa Maria Supersequence) of Southern Brazil

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From the Late Triassic Riograndia Assemblage Zone (Santa Maria Supersequence) of Southern Brazil Anais da Academia Brasileira de Ciências (2014) 86(4): 1673-1691 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 http://dx.doi.org/10.1590/0001-3765201420140455 www.scielo.br/aabc A new prozostrodontian cynodont (Therapsida) from the Late Triassic Riograndia Assemblage Zone (Santa Maria Supersequence) of Southern Brazil MARINA B. SOARES1, AGUSTÍN G. MARTINELLI1 and TÉO V. DE OLIVEIRA2 1Laboratório do Setor de Paleovertebrados, Departamento de Paleontologia e Estratigrafia, Instituto de Geociências, Universidade Federal do Rio Grande do Sul/UFRGS, Av. Bento Gonçalves, 9500, 90540-000 Porto Alegre, RS, Brasil 2Museu de Zoologia, Departamento de Ciências Biológicas, Universidade Estadual de Feira de Santana/UEFS, Av. Transnordestina, s/n, Novo Horizonte, 44036-900 Feira de Santana, BA, Brasil Manuscript received on September 8, 2014; accepted for publication on October 14, 2014 ABSTRACT We report here on a new prozostrodontian cynodont, Botucaraitherium belarminoi gen. et sp. nov., from the Late Triassic Riograndia Assemblage Zone (AZ) of the Candelária Sequence (Santa Maria Supersequence), collected in the Botucaraí Hill Site, Candelária Municipality, state of Rio Grande do Sul, Brazil. The new taxon is based on a single specimen (holotype MMACR-PV-003-T) which includes the left lower jaw, without postdentary bones, bearing the root of the last incisor, canine and four postcanines plus one partial crown inside the dentary, not erupted, and two maxillary fragments, one with a broken canine and another with one postcanine. The features of the lower jaw and lower/upper postcanines resemble those of the prozostrodontians Prozostrodon brasiliensis from the older Hyperodapedon AZ and Brasilodon quadrangularis and Brasilitherium riograndensis from the same Riograndia AZ. The inclusion of Botucaraitherium within a broad phylogenetic analysis, positioned it as a more derived taxon than tritylodontids, being the sister-taxon of Brasilodon, Brasilitherium plus Mammaliaformes. Although the new taxon is based on few cranial elements, it represents a additional faunal component of the Triassic Riograndia AZ of southern Brazil, in which small-sized derived non-mammaliaform cynodonts, closely related to the origin of mammaliaforms, were ecologically well succeed and taxonomically diverse. Key words: Cynodontia, Prozostrodontia, Santa Maria Supersequence, Triassic. INTRODUCTION based on correlations with Argentinean and The rich fossil tetrapod content from the Middle- Malagasy faunas (Abdala and Ribeiro 2010), Upper Triassic of southern Brazil is recorded in the non-mammaliaform cynodonts (Therapsida, beds related to the Santa Maria Supersequence, in Eucynodontia) account for a significant percentage which four recognized faunal tetrapod associations of the recorded taxa, taxonomically comprising succeed in time: Dinodontosaurus, Santacruzodon, the most diverse group. In fact, the diversity of the Hyperodapedon and Riograndia Assemblage Zones derived non-mammaliaform probainognathian (AZ) (Soares et al. 2011) (Fig. 1). In all of these cynodonts, grouped into the Clade Prozostrodontia bio stratigraphic unities, whose proposed ages are (sensu Liu and Olsen 2010), from the Upper Triassic Candelária Sequence (Santa Maria Supersequence) Correspondence to: Marina Bento Soares E-mail: [email protected] of southern Brazil is remarkable when compared to An Acad Bras Cienc (2014) 86 (4) 1674 MARINA B. SOARES, AGUSTÍN G. MARTINELLI and TÉO V. DE OLIVEIRA its global fossil record, based upon poorly preserved Bonaparte et al. (2003), and Minicynodon specimens. At least two prozostrodontians are Bonaparte et al. (2010). Riograndia is the most known from the Hyperodapedon AZ (Bonaparte and common cynodont of this AZ, represented Barberena 2001) and at least five from the younger by several exquisitely preserved specimens. Riograndia AZ (Bonaparte et al. 2001, 2003, 2005, Riograndia is considered as a basal member of 2010, 2012, Martinelli et al. 2005, Soares et al. 2011). the Tritheledontidae Clade (Bonaparte et al. 2001, Recent studies on these taxa have highlighted their Soares et al. 2011) or a basal Ictidosauria, sister- essential contribution to the understanding of the taxon of Tritheledontidae (sensu Martinelli et al. mammaliaforms origin (Bonaparte et al. 2005, Luo 2005, Martinelli and Rougier 2007). Irajatherium 2007, Abdala 2007, Liu and Olsen 2010, Bonaparte is still poorly represented in comparison to other 2012), illustrating the climbing mosaic of characters cynodonts from this AZ. However, its dentition developed during the Middle-Late Triassic transition resembles that of Chaliminia from the Late (Bonaparte et al. 2005, Martinelli and Rougier 2007, Triassic of Argentina and Pachygenelus from the Martinelli and Bonaparte 2011, Bonaparte 2012, Early Jurassic of Africa (Martinelli and Rougier Rodrigues et al. 2012, 2013). 2007). Hence, Irajatherium is positioned as a basal In the late Carnian Hyperodapedon AZ, Tritheledontidae (Martinelli et al. 2005, Martinelli the prozostrodontians Therioherpeton cargnini and Rougier 2007, Oliveira et al. 2011). Brasilodon, Bonaparte and Barberena (1975) and Prozostrodon Brasilitherium (Bonaparte et al. 2003, 2005, brasiliensis Bonaparte and Barberena (2001) are 2012), and Minicynodon (Bonaparte et al. 2010, recognized, both displaying several derived features 2012) are relatively well-documented and they are in the skull and dentition (e.g. lack of prefrontal crucial taxa in cynodont evolution because they and postorbital, constricted root in postcanines) constitute the sister-group of Mammaliaformes that positioned them as basal prozostrodontians (e.g., Bonaparte et al. 2003, 2005, 2012, Abdala (Liu and Olsen 2010). From this AZ, Charruodon 2007, Martinelli and Rougier 2007, Luo 2007, Liu tetracuspidatus Abdala and Ribeiro (2000) and and Olsen 2010, Rodrigues et al. 2012, 2013, Ruff Trucidocynodon riograndensis Oliveira et al. et al. 2014). Bonaparte et al. (2005) created the (2010) are also known. Charruodon was originally Family Brasilodontidae to include Brasilodon and related to Therioherpeton due to the morphology Brasilitherium, and, subsequently, other taxa were of the lower postcanines; nonetheless, based on included in it such as Protheriodon estudianti the deep and robust lower jaw (unexpected for (Bonaparte et al. 2006, Bonaparte 2012). Some Therioherpeton based on its skull), we consider that authors (e.g., Liu and Olsen 2010) pointed out the phylogenetic position of Charruodon cannot be that Brasilodon and Brasilitherium are synonym elucidated until new specimens come to light. On the and also that a monophyletic Brasilodontidae other hand, Trucidocynodon represents a member of has not been recovered (e.g., Abdala 2007). the Ecteniniidae (Oliveira et al. 2010, Martinez et al. Notwithstanding these issues that are currently 2013), which is positioned outside, as sister-group of under revision, the impact that the discovery of Prozostrodontia (Martinez et al. 2013). the brasilodontids from the Riograndia AZ of With regard to the prozostrodontian content, the southern Brazil has caused is unprecedented early Norian Riograndia AZ includes Riograndia (Bonaparte et al. 2003, 2005, 2012, Rodrigues et guaibensis Bonaparte et al. (2001) Irajatherium al. 2012, 2013). hernandezi Martinelli et al. (2005), Brasilodon In this contribution, we present a new pro- quadrangularis, Brasili therium riograndensis zostrodontian cynodont from the Riograndia AZ of An Acad Bras Cienc (2014) 86 (4) A NEW CYNODONT FROM THE LATE TRIASSIC OF BRAZIL 1675 the Candelária Municipality, state of Rio Grande with several features shared with Prozostrodon do Sul, Brazil. The new taxon is described, from the Hyperodapedon AZ and especially compared and included in a broad phylogenetic with Brasilodon and Brasilitherium from the analysis of cynodonts. Although based on a partial Riograndia AZ. The new specimen contributes lower jaw and maxillary fragments, the specimen to the understanding of the non-mammaliaform represents a new taxon of prozostrodontian prozostrodontian diversity. Fig. 1 - A: Geographical location of the Santa Maria Supersequence (Paraná Basin) in the state of Rio Grande do Sul, Brazil. B: Chronostratigraphy of the southern Brazilian Triassic, showing the four Assemblage Zones (AZ) based on tetrapods context. From Horn et al. 2004. Biostratigraphy after Soares et al. 2011. Abbreviations: Any, Anisian; Car, Carnian; Ind, Induan; Lad, Ladinian; Nor, Norian; Ole, Olenekian; Rha, Rhaetian. GEOLOGICAL SETTING AND BIOSTRATIGRAPHY and Scherer 2000, Zerfass et al. 2003). The Candelária Sequence encompasses the Hyperodapedon AZ The Upper Triassic Candelária Sequence corresponds (sensu Abdala et al. 2001), in which the most abundant to a third-order sequence placed in the Santa Maria components are the rhynchosaur Hyperodapedon Supersequence (sensu Zerfass et al. 2003, Horn et sp. and the traversodontid cynodont Exaeretodon al. 2014) (Fig. 1). The basal portion of the Sequence riograndensis Abdala et al. (2002). Besides the consists of a coarsening-upward succession that begins aforementioned cynodonts (e.g., Trucidocynodon, with red mudstones interbedded with small-scale Therioherpeton, Charruodon and Prozostrodon), its trough cross-bedded sandstone lenses. Rhytmites dinosaur content also deserves mention (Bittencourt and sigmoidal massive to climbing cross-laminated and Kellner 2009, Langer et al. 1999, Cabreira et al. sandstone bodies are also present
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